Number 336: 1-12

ISSN 1026-051X

June 2017http/urn:lsid:zoobank.org:pub:C7175324-05B5-494A-8973-7AEF4E641B07

PALAEARCTIC SPECIES OF THE GENUSLAMPROCHROMUS MIK, 1878 (DIPTERA:DOLICHOPODIDAE)
I. Ya. Grichanov1,*), Azam Ahmadi2)
1) All-Russian Institute of Plant Protection, Podbelskogo 3, St. PetersburgPushkin, 189620, Russia. * Corresponding author, E-mail: grichanov@mail.ru
2) Baran Plant Protection Institute, Arak, Markazi Province, Iran.
A new long-legged fly species Lamprochromus occidasiaticus sp. n. from Iranand Turkey is described and illustrated. The new species appears to be quite distinct
in having elongate-ovate postpedicel of antenna, abdomen entirely dark, hind coxa
and femur partly dark, and male phallus with strongly curved thick seta. A catalog
and key to ten Lamprochromus species inhabiting the Palaearctic Region isprovided.
KEY WORDS: Dolichopodidae, Lamprochromus, new species, key, checklist,
Iran, Turkey, Palaearctic Region.
И. Я. Гричанов*, Азам Ахмади. Палеарктические виды рода Lamprochromus Mik, 1878 (Diptera: Dolichopodidae) // Дальневосточный энтомолог.
2017. N 336. С. 1-12.
Из Ирана и Турции описан Lamprochromus occidasiaticus sp. n. Новый видотличается от других видов рода удлиненно-овальным третьим члеником
усика, полностью темным брюшком, темными задними тазиками и частично
темными задними бедрами, сильно изогнутой толстой щетинкой на фаллусе
самца. Даны каталог и определительная таблица десяти видов рода Lamprochromus Палеарктики.
1

*Корреспондирующий автор, Всероссийский научно-исследовательский
институт защиты растений, Санкт-Петербург, Пушкин, Россия.
INTRODUCTION
There are 13 Holarctic Lamprochromus species including 9 from the Palaearctic
Region and one from the Afrotropical Region (Grichanov, 2014). The genus
belongs to the subfamily Sympycninae, differing from other genera by thefollowing combination of characters. Body small, often yellow-brown to bluishblack; antennal arista-like stylus dorsal or basodorsal; mesonotum with two velvety
black or matt brown large lateral spots above the wing base, sometimes poorly
developed; 3 or 4 pairs of dorsocentral bristles; acrostichal setae in two regularrows; hind femur with true subapical bristle (Grichanov et al., 2011). A key to the
Palaearctic species was provided by Negrobov & Chalaya (1988).
Treating a new material collected during 2016 season from 64 sites in the Markazi
and Lorestan provinces, we have found a male and a female of a new peculiar
Lamprochromus species described below. Additional male and female of the same
species have been found in the Turkish collection of the Zoological Museum of
Moscow State University, Moscow, Russia. In this paper a catalog and identification key to Palaearctic species of Lamprochromus are also presented.
MATERIAL AND METHODSThe holotype and paratypes of the new species have been sampled by standard
sweep net and are housed at the Zoological Institute of the Russian Academy of
Sciences, St. Petersburg (ZIN) and the Zoological Museum of Moscow State
University, Moscow (ZMUM). The other material examined is deposited also at the
Department of Zoology, Tel Aviv University, Israel (TAU).
The specimens were studied and illustrated with ZEISS Discovery V-12 stereomicroscope and AxioCam MRc5 camera. Morphological terminology follows
Cumming and Wood (2009). The relative lengths of the podomeres are given in
millimeteres. Body length is measured from the base of the antenna to the tip ofabdominal segment 7. Wing length is measured from the base to the wing apex.
Figure showing the male genitalia in lateral view is oriented as they appear on the
intact specimen (rotated 180° and lateroflexed to the right), with the morphologicallyventral surface of the genitalia facing up, dorsal surface down, anterior end facing
right and posterior end facing left.
TAXONOMYFamily Dolichopodidae
Genus Lamprochromus Mik, 1878
Lamprochromus Mik, 1878: 4 (subfam. Sympycninae).

Type species: Chrysotus elegans Meigen, 1830 [=Lamprochromus bifasciatus(Macquart)], by original designation.
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Key to the Palaearctic species of the genus Lamprochromus
1. Males: 2nd and 3rd abdominal tergites usually yellow, transparent .................... 2– Females: abdomen usually entirely dark metallic green ..................................... 10
2. Abdomen entirely dark metallic green ................................................................. 3– Abdomen yellow at base ....................................................................................... 5
3. Postpedicel triangular, pointed at apex, slightly longer than high; coxae yellow,
mid coxa with dark stripe; 1.7 ............................................... L. defectivus Strobl– Postpedicel ovate or subtriangular, rounded at apex, at least 1.5 times longer thanhigh; mid and hind coxae largely dark ............................................................... 4
4. Postpedicel 1.5 times longer than high; fore coxa covered with long fine anteriorcilia; mid and hind coxae yellow, blackish at base; hind femur entirely yellow;
cercus as long as epandrium; 2.0 ......................................... L. dalmaticus Parent– Postpedicel 2 times longer than high; fore coxa covered with short light anterior
hairs; mid and hind coxae brown, at most yellow at apex; hind femur brown orbrownish in distal half or third at least dorsally; cercus distinctly (3/5) shorter
than epandrium; 2.3–2.5 .................................................. L. occidasiaticus sp. n.
5. Postpedicel triangular, pointed at apex, with long sparse hairs; 1.5–2.0 ...............
....................................................................................... L. bifasciatus (Macquart)– Postpedicel ovate-triangular, rounded at apex, usually with short dense hairs ..... 6
6. Postpedicel not longer than high; at least scape and pedicel of antenna yellow;
each lobe of phallus with long pointed process apically .................................... 7– Postpedicel distinctly longer than high; antenna entirely brown or black; phallus
with strong spines or without spines apically .................................................... 8
7. Fore coxa with yellow bristles at apex; hind tibia flattened laterally, mainly brown;
1.5–2.0 .................................................................................. L. speciosus (Loew)– Fore coxa with dark bristles at apex; hind tibia simple, mainly yellow; 1.6 ..........
.......................................................................... L. kowarzi Negrobov et Tshalaja
8. Phallus without spines apically; surstylus rounded at apex; frons violet-green with
red shine; 1.2 ............................................ L. moraviensis Negrobov et Tshalaja–Each lobe of phallus with strong spine apically; surstylus truncated or angular at
apex; frons green, with violet shine in middle ................................................... 9
9. Face silvery grey; mesonotum with poorly developed brown lateral spots; 2.1 ....
.................................................................. L. buchtojarovi Negrobov et Tshalaja– Face silvery white; mesonotum with velvety black or brown lateral spots; 2.0 .....
............................................................................................ L. semiflavus (Strobl)
10. Abdomen yellow at base .................................................................................. 11– Abdomen entirely dark metallic green ................................................................ 13
11. Fore coxa with yellow bristles at apex ................................. L. speciosus (Loew)– Fore coxa with dark bristles at apex. ................................................................... 12
12. Pleura black-green ................................................................... L. amabilis Parent– Pleura largely yellow-brown ............................... L. kowarzi Negrobov et Tshalaja
13. Postpedicel almost oval, rounded at apex .......................... L. semiflavus (Strobl)– Postpedicel triangular, angular at apex ............................................................... 143

14. Hind femur brown in distal half or third at least dorsally .....................................
.......................................................................................... L. occidasiaticus sp. n.
– Hind femur entirely yellow ..................................................................................15
15. Mid coxa entirely yellow; hind basitarsus light at base; wing anal vein distinct ..
....................................................................................... L. bifasciatus (Macquart)– Mid coxa with grey spot; hind basitarsus entirely dark; wing anal vein absent .......
................................................................................................ L. defectivus Strobl
Lamprochromus occidasiaticus Grichanov et Ahmadi, sp. n.
Figs 1–6
TYPE MATERIAL. Holotype – ♂, Turkey: near Manavgat River, 29.III 2007,
N. Vikhrev [ZMUM]. Paratypes: 1 ♀, same label [ZMUM]; 1 ♂, 1 ♀, Iran: Markazi
province: Seyed Esmaeel road, 22.V 2016, 34°13.68' N, 49°08.77' E, 2073 m, A.
Ahmadi [ZIN].
DESCRIPTION. Male (Figs 1–5). Head: frons shining blue-violet; face black,densely silvery-white pollinose, gradually narrowed downward; face under antennae
nearly 2 times as wide as height of postpedicel; eyes widely separated in lower halfof face; palpi and proboscis black with short black setae; antenna black; scape bare;pedicel simple, with ring of short setulae and 1–2 longer setae dorsally; postpedicel
elongate ovate, swollen at base, with rounded apex, 2 times as long as high, densely
short pubescent; stylus simple, shortly pubescent, basodorsal, 1st segment of stylus
thick; postoculars in single row, upper setae black, lower postoculars white. Length
ratio of scape to pedicel to postpedicel to stylus (1st and 2nd segments), 0.1: 0.06:
0.2: 0.06: 0.56.
Thorax: mostly bluish or greenish black, grey pollinose; mesonotum anteriorly
with pair of shining violet-black spots; notopleura with large mat-black spot; setae
black; proepisternum without strong setae, with 2–3 short cilia; 4 (2+2)
dorsocentrals of approximately equal length with 2–3 hairs in front of the 1st seta;
about 10 pairs of biseriate acrostichals; 1 pair of strong scutellar setae and pair oflateral hairs.
Legs: including fore coxa mostly yellow; mid and hind coxae brown, at mostyellow at apex; tarsi brown or black from apex of basitarsus; fore and mid femora
yellow or brownish dorsally, hind femur brown dorsally in apical half to third. Forecoxa covered with short light anterior hairs, with some strong black apical setae,
mid coxa with few black setae, and hind coxa with 1 black seta. Fore femur with 1
posteroventral subapical stiff cilia; fore tibia with 1 anterodorsal seta at 1/4, withweak anterodorsal serration along distal 2/3; fore tarsus simple. Length ratio of fore
tibia to tarsus (segments from first to fifth), 0.62: 0.31: 0.14: 0.10: 0.09: 0.09. Mid
femur with 1 anterior and 1 posterior subapical setae; mid tibia with anterodorsal
bristles at 1/4 and 1/2, with 2 shorter posterodorsal setae and 4-5 apical setae; midtarsus simple. Length ratio of mid tibia to tarsus (segments from first to fifth), 0.86:
0.42: 0.21: 0.17: 0.11: 0.11. Hind femur with anterior subapical seta, with weak subapical posteroventral seta; hind tibia with 2 strong anterodorsal, 2–4 posterodorsal,
3–5 inconspicuous ventral and 4–5 apical setae. Length ratio of hind tibia to tarsus(segments from first to fifth), 1.09: 0.31: 0.34: 0.22: 0.16: 0.14.
4Wing: greyish with brown veins; ratio of cross-vein dm-cu to apical part ofCuA1, 14: 58; distal part of M1+2 weakly convex anteriorly, almost parallel toR4+5; anal vein indistinct, fold-like; lower calypter yellow with white setae; halter
yellow.

Figs. 1–5. Lamprochromus occidasiaticus, sp. n. 1 – male habitus; 2 – male head, lateralview; 3 – male wing; 4 – male hypopygium, lateral view, with inset showing phallus; 5 –male hypopygium, ventral view.

Abdomen: metallic bluish or greenish-black, with black vestiture; hypopygium
brown-black with black cerci. Epandrium rounded-oval (lateral view). Epandrial
lobe as narrow strip along ventral margin of epandrium extending from base of
hypandrium to base of surstylus, with short seta at middle and 2 short setae at apex.
Hypandrium fused with epandrium at base, with short and broad semi-oval medialplate and 2 long and narrow symmetrical lateral arms reaching base of surstylus.
Parameral sheat sclerotised, widely bifurcated distally, with narrow symmetricallateral arms. Phallus of typical shape for genus (see Negrobov & Tshalaja, 1988),
gradually expanded from base of hypandrium distally (lateral and ventral view),
bilobed apically; each lateral lobe angular ventrally at apex, bearing long narrowdorsal process and strongly curved thick apical spine. Surstylus with fused lobes,
5forming broad plate, ovate or subtriangular (lateral view), with strongly curved
pointed apex (ventral view), covered with numerous setae, which rather long at
middle of surstylus ventrally. Cercus subtriangular, distinctly shorter than epandrium(3/5), with long black setae; cercus nearly 2 times longer than wide at middle, withacute distal angle and rounded lateroventral angle; cerci fused at base, forming
unpaired projected organ.
Female (Fig. 6): Antenna shorter, postpedicel as long as high, Face wider thanin male, 2 times wider than postpedicel, nearly parallel sided. Palpus larger subtriangular. Length ratio of fore tibia to tarsus (segments from first to fifth), 70: 31:
13: 9: 8: 9. Length ratio of mid tibia to tarsus (segments from first to fifth), 90: 44:
18: 15: 8: 10. Length ratio of hind tibia to tarsus (segments from first to fifth), 120:
32: 35: 23: 14: 12.

Fig. 6. Lamprochromus occidasiaticus sp. n., female habitus.

MEASUREMENTS (in mm). Body length without antennae: ♂ 2.5, ♀ 2.3;
antenna length: ♂ 0.8, ♀ 0.7; wing length ♂ 2.4, ♀ 2.2; wing width 0.8.
DISTRIBUTION. Iran, Turkey.
ETYMOLOGY. The species name originates from Latin “Asia occidentalis”(West Asia).
6

DIAGNOSIS. The new species is close to Lamprochromus dalmaticus Parent,
differing in longer postpedicel, 2 times longer than high; mesonotum having anteriorly a pair of shining violet-black spots; wing anal vein being indistinct, fold-like;fore coxa covered with short light anterior hairs; mid and hind coxae brown, at mostyellow at apex; brown or brownish hind femur in distal half or third at least
dorsally; cercus distinctly (3/5) shorter than epandrium. L. dalmaticus was describedwith postpedicel 1.5 times longer than high; mesonotum having anteriorly a pair ofcopper spots; wing anal vein being rather distinct; fore coxa covered with long fineanterior cilia; mid and hind coxae yellow, blackish at base; entirely yellow hindfemur; cercus as long as hypopygium. In addition, Lamprochromus occidasiaticus
is distinctly larger, than L. dalmaticus (see key above).
Catalog of Palaearctic Lamprochromus species and new records
Lamprochromus amabilis Parent, 1944
Lamprochromus amabilis Parent, 1944: 122. Type locality: China: “Chensi, Yulinnfou”.

DISTRIBUTION. China (Shaanxi).
NOTES. The species was described by a single female holotype and was neverrecorded again.
Lamprochromus bifasciatus (Macquart, 1827)
Medetera bifasciata Macquart, 1827: 48; Zetterstedt, 1843: 463 (as synonym of
Rhaphium fasciatum Meigen, 1824); Parent, 1938: 661 (as synonym of Lamprochromus
elegans); Meuffels et al., 1991: 100 (Lamprochromus); Grichanov et al., 2011a: 35, Fig. 55(Lamprochromus); Grichanov et al., 2011b: 32, Fig. 70 (Lamprochromus). Type locality: notgiven [North France].
Chrysotus elegans Meigen, 1830: 362; Loew, 1857: 17 (Gymnopternus); Kowarz, 1868:
219 (Sympycnus); Mik, 1878: 4 (Lamprochromus); Pârvu, 1984: 202, Fig. 10 (Lamprochromus); Parent, 1938: 661, Figs. 914–916 (Lamprochromus); Negrobov & Tshalaja, 1988:
898, Figs. 4–5 (Lamprochromus). (Synonymised by Meuffels et al., 1991: 100). Type
locality: not given.
Medetera gratiosa Meigen, 1838: 158; Parent, 1925b: 54, 57 (as synonym of
Lamprochromus elegans).
Dolichopus bifasciellus Zetterstedt, 1843: 608; Loew, 1857: 44 (Sympycnus); Mik, 1878:
7 (Lamprochromus). (Synonymised by Kowarz, 1868: 219). Type locality: “Scania, Esperöd,
Ostrogothia, Häradshammar, Lärketorp, Wadstena, Uplandia, Holmiam, Lapponia Umensis,
Wilhelmina, in alpe Dovre, Norvegiae” [Sweden, Norway].
Medetera cingulum Parent (Meigen in coll.), 1925a: 137 (footnote); Parent, 1925b: 52,
57 (as synonym of Lamprochromus elegans); Parent, 1938: 661 (as synonym of Lamprochromus elegans). Type locality: not given.

DISTRIBUTION. Austria, Belgium, Bulgaria, Czech, France, Germany, Hungary, Italy, Netherlands, Norway, Poland, Romania, Russia (Leningrad), Slovakia,
Spain, Sweden, Switzerland, UK. Here excluded from the fauna of Israel.
7

Lamprochromus buchtojarovi Negrobov et Tshalaja, 1988
Lamprochromus buchtojarovi Negrobov & Tshalaja, 1988: 797, Figs. 7–9. Type locality:
Russia: Chuvashia, Cheboksary env.

DISTRIBUTION. Russia (Cheboksary).
Lamprochromus dalmaticus Parent, 1927
Lamprochromus dalmaticus Parent, 1927: 45. Type locality: Croatia: “Côte Dalmate,
Raguse”.

DISTRIBUTION. Croatia.
NOTES. A record of this species from Erzurum, Turkey (Naglis, 2011) belongs
to L. occidasiaticus (S. Naglis, 2016 pers. comm.; unreferenced).
Lamprochromus defectivus Strobl, 1899
Lamprochromus defectivus Strobl, 1899: 121. Type locality: Spain: Algeciras.

DISTRIBUTION. Greece (Crete) [needs confirmation], Spain.
NOTES. Negrobov & Tshalaja (1988) described L. defectivus male postpedicel
with pointed apex (in key), but erroneously figured the postpedicel with rounded
apex (Negrobov & Tshalaja, 1988: Fig. 12).
Lamprochromus kowarzi Negrobov et Tshalaja, 1988
Lamprochromus kowarzi Negrobov & Tshalaja, 1988: 797, Figs. 4–6; Grichanov, 2007a:
67 (as Lamprochromus bifasciatus); Grichanov, 2007b: 146 (as Lamprochromus speciosus,
in part); Grichanov & Tomkovich, 2009: 107 (as Lamprochromus bifasciatus); Grichanov &Ovsyannikova, 2016: 288 (as Lamprochromus speciosus). Type locality: Slovakia: Lozonez.

MATERIAL EXAMINED. Azerbaijan: Lәnkәran, Hirkan-Bürcәli env., 16, 17,
18.V 2009, 1 ♂, 2 ♀, Grichanov & Tomkovich [ZMUM]; Israel: Rosh Ha’Ayin,
10.IV 1976, 14 ♂♀, A. Freidberg [TAU]; Golan [Heights], Qusbiye, 15.IV 1982,
1♂, F. Kaplan [TAU]; Park HaYarden, 30.IX 1982, 1 ♀, A. Freidberg [TAU];
Nahal Tut, 18.V 1982, 1 ♀, A. Freidberg [TAU]; Russia: Chechnya, Sernovodskaya, health resort, 43.33°N, 45.17°E, 15.VI 2016, 1 ♀, Grichanov [ZIN].
DISTRIBUTION. Azerbaijan, Israel, Russia (Chechnya), Slovakia.
Lamprochromus moraviensis Negrobov et Tshalaja, 1988
Lamprochromus moraviensis Negrobov & Tshalaja, 1988: 795, Figs. 1–3. Type locality:
Moravia S.

DISTRIBUTION. Czech Republic.
8

Lamprochromus semiflavus (Strobl, 1880)
Diaphorus semiflavus Strobl, 1880: 58; Mik, 1881: 346 (as synonym of Lamprochromus
bifasciatus); Parent, 1925: 138 (as Lamprochromus strobli nom. nov.); Negrobov, 1991: 58(as synonym of Lamprochromus elegans). Type localitty: Austria: Conventgarten.
Lamprochromus strobli Parent, 1925a: 141 (unnecessary nom. nov. for Lamprochromus
semiflavus); Parent, 1938: 663, Figs. 920–921; Negrobov & Tshalaja, 1988: 898, Figs. 6–8.

MATERIAL EXAMINED. Russia: Voronezh Region, Liski distr., Divnogorye,
8-9.VII 1994, 8 ♂♀, Zlobin [ZIN]; Pskov Region, Velikie Luki env., 7, 10, 26, 29,
30.VI 1997, 10 ♂♀, Grichanov [ZIN]; Orel city, Orlik River bank, 6.VII 2015, 1 ♀,Grichanov [ZIN]; Moscow region, Naro-Fominsk, park, 55.39 N, 36.74 E, 160 m,
22.VI 2013, 1 ♂, D. Gavryushin [ZMUM].
DISTRIBUTION. Austria, Belgium, Bulgaria, Czech, Finland, France, Germany,
Hungary, Netherlands, Poland, Portugal, Russia (Orel, Pskov, Voronezh), Turkey(Adiyaman), UK.
NOTES. Soon after description, Diaphorus semiflavus was placed in synonymy
with Lamprochromus elegans (now synonym of L. bifasciatus), single species of
the genus known at that time (Mik, 1881). Later, the synonymy was republished byBecker (1918) and Negrobov (1991) without explanations. Strobl (1899) described
L. defectivus, distinguishing his new species from the misidentified L. elegans bypointed, rather than rounded, apex of postpedicel. Parent (1925a), revising available
collections and original descriptions, found that both L. defectivus and L. elegans
well differ from L. semiflavus and L. speciosus by pointed, rather than rounded,
apex of postpedicel. He restored virtually the name “semiflavus”, but (having notypes in his hands) he proposed a new name L. strobli for the species concept
published by Strobl under the name D. semiflavus. Parent (1938) mentioned thelatter name in synonymy to L. strobli. The recent catalogs and checklists follow
synonymy of either Parent or Mik and Negrobov. Here we consider the name“semiflavus” as valid name, the opinion of Parent as reasonable, and we place thename L. strobli in synonymy to L. semiflavus according with the principle ofpriority. The hypopygium including phallus of L. semiflavus is identical to that of L.
buchtojarovi as pictured by Negrobov & Tshalaja (1988).
Lamprochromus speciosus (Loew, 1871)
Sympycnus speciosus Loew, 1871: 299; Kowarz, 1889: 175 (Lamprochromus); Parent,
1938: 662, Figs. 917–919 (Lamprochromus); Pârvu, 1984: 202, Fig. 11 (Lamprochromus);Negrobov & Tshalaja, 1988: 798, Figs. 1–3 (Lamprochromus); Grichanov, 2007b: 146(Lamprochromus speciosus, in part). Type locality: Tajikistan: “Sarawschan Thal” [=Zeravshan valley].

MATERIAL EXAMINED. South Tajikistan: Dusti env., 24,VIII.1987, 1 ♂, Grichanov; Israel: Park HaYarden, 30.IX.1982, 4 ♂♀, A. Freidberg [TAU]; Israel: Ein
Hajla, 11.V.1977, 3 ♂♀, A. Freidberg [TAU].
9DISTRIBUTION. Bulgaria, Egypt, France, Greece, Hungary, Iran, Iraq, Israel,
Romania, Russia (Krasnodar), Spain (Canary Is.), Uzbekistan, Tajikistan, Turkey(Muğla), Ukraine (Odessa). Here excluded from the fauna of Azerbaijan and Chechen
Republic of Russia.
Discussion
Species of the genus Lamprochromus are rather rare in collections. Six of thenine previously described Palaearctic species have been known only from their
original descriptions, often from single type specimen. Our study of series of L.
kowarzi, L. semiflavus and L. speciosus males and females has proved the stabilityof such species-specific characters as colouration of coxae and femora, length ofantennal postpedicel and length of male cercus. We had no opportunity to examine
the types or topotypes of L. dalmaticus; nevertheless, we believe that the complex
of diagnostic characters in L. occidasiaticus sp. n. (see diagnosis of the new species)
clearly distinguishes the latter from the L. dalmaticus despite possible intraspecificvariation of some characters.
Generally, the Lamprochromus adult habitus and morphology of hypopygium
correspond to the concept of the subfamily Sympycninae. Germann et al. (2011)
have analysed the phylogeny of Dolichopodidae and suggested that the genus Lamprochromus is positioned within the Rhaphiinae clade. However, the latter subfamily
still deserves a more detailed study, being at present polyphyletic assemblage ofgenera. L. bifasciatus is a typical wetland species in reed ditches and permanent
reed marshes of Belgium (Decleer et al. 2015). The West Palaearctic species of theLamprochromus occur in the countries with relatively mild climate, from southern
Scandinavia in the North, to North Africa and Central Asia in the South.
ACKNOWLEDGEMENTS
The authors are sincerely grateful to Drs. Andrei Ozerov and Nikita Vikhrev(Moscow, Russia) and to Dr. Amnon Freidberg (Tel Aviv, Israel) for their kindness
in providing specimens for study. Dr. Stefan Naglis (Zurich, Switzerland) andanonymous reviewers kindly commented on earlier drafts of the manuscript.
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